A Primer for the Exercise and Nutrition Sciences

Thermodynamics, Bioenergetics, Metabolism

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Christopher B. Scott, PhD
Gorham ME
USA

ISBN 978-1-60327-382-4 e-ISBN 978-1-60327-383-1

DOI 10.1007/978-1-60327-383-1
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Preface

What a journey writing this text has been. The lengthy voyage started well before
the idea hatched of authoring a text that contained the word “thermodynamics”! I
was informed by my good friend and sometimes colleague Dr. Jose Antonio that by
including that word in the title, nutritionists and exercise physiologists might avoid
the subject. But almost every step of my expedition was taken on a rather solid
foundation of thermodynamics and as such the topic could not possibly be omitted
from the title or the text of a book about bioenergetics and energy expenditure.
I am not a physicist. In fact I first went to college to become a football coach.
That vocational choice began to deteriorate when taking the mandatory anatomy and
physiology courses required of all physical education majors. This information was
exciting; my interest in physical education began to wane. During sophomore year,
I answered an advertisement in the school newspaper requesting research subjects.
The request was made by a master’s student who was correlating the presence of
the anaerobic threshold with the rating of perceived exertion (RPE) during treadmill
running to exhaustion (it happens at the point when the subject perceived that the
work being performed was “somewhat hard” to “hard”). This was a cool endeavor!
So cool that I asked the study’s author if it were possible to hang out and help. Soon,
I was calibrating the equipment in addition to helping with subject testing. It was
this experience more than anything else that defined my chosen career path.
During my senior year internship, I met Dr. La Von Johnson, who offered a full
scholarship to obtain a master’s degree in sports medicine. Dr. Johnson remains the
most influential of all my academic acquaintances; a true teacher, professor, mentor,
and friend. Our mutual academic interest in strength, speed, and power planted a
seed that still grows today (thank you, Dr. Johnson!).
With graduate school supposedly complete, I entered the workforce; a manager
or exercise programmer for a fitness center. It soon became clear that this was not go-
ing to be a career. Later, at yet another fitness facility, more and more time was spent
in academic libraries doing “armchair research.” A short year-long stint with an au-
thor who was writing fitness and nutrition-related material soon followed. As a re-
search editor, I found myself spending even more time in armchair research-related
endeavors. My future was seemingly in view: Do not just read about research, do

v
vi Preface

it. The next step, it seemed, was to become something of a respected authority in
the exercise sciences. Becoming a Ph.D. scientist should have something to do with
that! A doctoral degree was needed to help accomplish this. Surprisingly, every
school I applied to rejected my admission requests. Although painful at the moment,
these academic institutions were correct in doing so. The educational background
I had chosen emphasized exercise as a human behavior, motivating or instructing
others on how to properly train. I was attempting to get into Ph.D. programs that
were built on the more basic sciences, where exercise was used as a model or tool
to study human physiology. Simply put, I was not adequately trained to become an
authority in exercise science.
I was not alone on the preliminary path I had chosen. Now as a college professor,
students routinely enter my office with an agenda of avoiding academic challenge,
choosing classes that ignore a second semester of chemistry, a full year of organic
chemistry and not even considering biochemistry, molecular biology, and cell phys-
iology. A desperation phone call to a world-renowned exercise physiologist, whom
I had never met, helped me through my “why didn’t I apply myself” crisis. As an
armchair researcher, I had read many of his professional publications and was quite
shocked that he took my phone call. Dr. Phil Gollnick (1935–1991) convinced me
to go back to school to get another master’s degree, this time with an emphasis on
science and research. And so at the age of 27 I did.
Dr. Fred Roby served as my next graduate school mentor and with his guid-
ance my thesis was completed and published (“The maximally accumulated oxygen
deficit as an indicator of anaerobic capacity”). In the academic arena, while most
graduate students are doing work dictated by laboratory heads and mentors, Dr.
Roby allowed me to pursue a project of my own interest. His encouragement is
appreciated (and I will never forget his ever-present smile).
My next career move seemed somewhat of a miracle. Years earlier, an internship
was completed at the Cooper Institute for Aerobics Research in Dallas, TX, under
the supervision of Dr. Jill Upton (among others). Dr. Upton recognized my passion
for exercise science and provided my superiors with glowing reviews. Though the
internship itself was brief (∼6 weeks), friends and colleagues were made, and I took
it upon myself to visit the Aerobics Institute whenever traveling through Dallas.
How shocked I was to receive a phone call that informed me of a position opening
up at the Cooper’s Aerobics Institute. I applied, was accepted, and immediately took
the job as a research associate in charge of the exercise physiology laboratory. This
was an incredible experience. My colleagues at the Cooper Institute for Aerobics
Research are too numerous to name, but working with the likes of Dr. Neil Gordon,
Dr. John Duncan, Kia Vaandrager, Dr. Steve Blair, and Dr. Bill Kohl was a dream
come true. I thank them for providing me not only with access to their expertise, but
allowing me to collect data, be a part of research design and in the process become
an author and colleague on well over a dozen scientific manuscripts. Moreover, this
exercise physiology laboratory was and is world renowned to the higher echelon
of accomplished athletes. The list of elite world-class amateurs and professionals
that came to the lab for testing was nothing less than astounding, and I was the
guy who tested them. Every exercise stress test I had performed at the Aerobic
Preface vii

Institute included the measurement of gas exchange (along with an ECG). Even
so, my passion for anaerobic energy expenditure, which fueled strength, speed, and
power, never faltered (a colleague once informed me, “This is the Cooper Institute
for Aerobic Research, not Anaerobic Research”).
A physician at the Cooper Institute had left to join a private cardiology practice,
I was asked to come along. During the ensuing interview I was questioned, “How
would you fit into our facility? Your background is in the testing of athletes not
diseased patients.” My reply was immediate and straight forward, “been there done
that, it’s now time to apply my exercise stress testing skills to cardiac and pulmonary
patients”. And so I did, for another 5 years (in two cardiology practices). This ex-
perience too was rewarding. Patients who could not walk for 2 min on the treadmill
were tested (this was a far cry from athletes who could run for almost 30 min). But
the times were soon to be changing. Business-related cuts being made in “our” prac-
tice were harsh and I felt that, void of an MD degree, my head was on the employee
termination chopping block. It was time to leave medicine, “call my own shots,” and
to focus exclusively on anaerobic and aerobic metabolism in the pursuit of a doctoral
degree. During the ensuing good-bye handshakes, I inquired about how many exer-
cise stress tests I had performed. The answer was, “somewhere between 5,000 and
10,000 tests;” again, all of them completed with full gas exchange measurements
and ECG interpretation.
In hindsight I had to this point undergone an extensive 10-year apprenticeship.
Academic texts were not being memorized; exercise physiology was actively be-
ing practiced. Taking objective notice of my interests, it became clear that I was
spending more time reading comparative physiology studies, where animals were
being used as exercise models, not athletes or patients. This was a new frontier
for me and I pursued it. But before moving on, notes were beginning to be col-
lected that eventually became part of the text you are now reading. Such note taking
was encouraged by several encounters I had with Dr. Roland Coulson (1915–2004).
Since graduate school, my interest has never waned in the measurement of anaero-
bic metabolism and it was Dr. Coulson’s metabolic studies using alligators that drew
me toward him (reptile muscles are fueled in large degree by anaerobic metabolism
during exercise). The elderly but very active Dr. Coulson was no longer accepting
Ph.D. students when I first contacted him, but he was always willing and able to
talk about metabolism. It was he who informed me, “if you want to really learn
something, write a book about it” (he also was the first authority figure with whom
I had a scholarly discussion on the metabolism of dinosaurs; I was in heaven that
night). Dr. Coulson’s statement represents the very start of my foray into thermody-
namic studies. Visits to Roland Coulson were an experience where as a guest at his
New Orlean’s residence I was awoken at around 3:00 in the morning, accompanied
by him on bicycle to his laboratory, and then promptly helped clean alligator poop
from his numerous cages (he was conducting feeding and growth studies)!
Toward the end of my clinical career I had begun researching Ph.D. programs
that focused on both zoology and physiology and found one that accepted me. I
was now 37-years old. Obtaining this degree involved difficulties, as most change
does. I put my house on the market (in Savannah, Georgia) and moved West (to
viii Preface

Wyoming). My salary fell from $45,000/yearly to an $8,500/yearly stipend. At first

I worked with an animal model that underwent seasonal hibernation (better referred
to as metabolic torpor). My friends questioned the move (“What does this have to do
with exercise physiology?”), but I had a rationale. Metabolic torpor is the exact op-
posite of VO2 max; it represents the lowest possible limits of aerobic metabolic en-
ergy expenditure, not the highest. I wanted to understand both ends of the metabolic
spectrum. Sadly, this experience did not last, after a year and a half my doctoral
advisor canned our project and just as I was planning to abandon the program I was
approached (and “saved”) by Dr. Paul Thomas and Dr. Richard McCormick who
needed assistance with a cardiology-related project. I accepted. The experience was
again rewarding, spending several years at the cellular and molecular level, examin-
ing a rat heart’s re-modeling of connective tissue after a myocardial infarction. The
assistance of Drs. Scott Boitano, Robert George, and Paul Wade will not be forgot-
ten (they recognized my deficiency in cellular course work and designed a program
to eliminate this flaw; I now present aspects of cellular physiology as part of most
every undergraduate course I teach).
During my doctoral studies notes for this text continued to be collected, ending
at something around 400 pages. The experience was a resounding success. But even
so, while I had much material, the project was nothing more than an amateurish col-
lection of scribblings. It took another several years to focus these pages toward the
subject of whole-body energy exchange. The first accomplishment was to write the
thermodynamics section. Thanks go to Dr. Jerry Bell (of the American Chemical
Society), who faithfully and promptly answered my many thermodynamic queries.
Gratitude also is given to Dr. Zoran M. Djurisic, a combustion scientist at U Califor-
nia, Berkeley, who worked with me in detailing the molecular differences in glucose
and fat combustion. (Apologies to Dr. Bell and Dr. Djurisic if I take too much liberty
with thermodynamic interpretation.)
As I explored potential publishers for this text I was somewhat taken back by
the reply of some editor’s, “We already have an extensive collection of biochem-
istry texts, none more are needed.” I had not explained myself well. It was not a
biochemistry textbook I was thinking about, though it certainly did employ some
biochemistry. Moreover, this text was not an attempt to re-name exercise biochem-
istry or bioenergetics as other have. Bioenergetics is presented here not as a series
of independent biochemical reactions that supply ATP to working muscle, but as a
mechanical energy-exchange device of sorts with moving parts and all; as much en-
gineering as biochemistry. The last section takes thermodynamic and bioenergetic
principles and applies them toward the estimation of metabolic energy expenditure.
Many of these applications are new. In fact, I have been told more than once that
some of the ideas presented represent little more than “excrement” on paper. Even
so, no critic has ever satisfactorily answered fundamental questions I have had for
decades and attempt to answer here (a typical response to my queries was, “you need
to read more”). Some examples include: How and why does a measurement of oxy-
gen uptake represent the energy expenditure of a metabolism that does not utilize
oxygen? Incorrect answer: because it does. Correct Answer: it does not. Why is the
energy associated with the oxidation of glucose greater than that of fat per volume
Preface ix

of oxygen consumed? Incorrect answer: glucose is a more efficient fuel than fat.
Correct answer: because glucose metabolism contains an anaerobic component, fat
does not. Why is metabolic biochemistry based on a diffusion-oriented system when
evidence indicates it cannot possibly be? Incorrect answer: because that is the way
it has been. Correct answer: it should not be.
My passion for the measurement and estimation of both aerobic and anaerobic
energy expenditure during strength, speed, and power has never diminished and it is
in this area that I continue much of my research. I also am very much interested in
markers of aerobic and anaerobic energy exchange in the diagnosis and prognosis
of heart vs. lung vs. skeletal muscle limitations to exhaustive exercise.
Wrapping this preface up in full circle, in a long-ago conversation with Dr. Jose
Antonio (about 15 years prior), I emotionally laid out a case as to why the oxygen
consumed in the recovery from exercise could not represent anaerobic energy ex-
penditure during the exercise. Joey informed me, “there are more people than you
think Chris, who will not disagree with you.” I hope my case, more thoroughly dis-
played in these pages, provides some meaningful answers.

Gorham, ME Christopher B. Scott

Contents

1 Introduction: Thermodynamics, Bioenergetics, Metabolism . . . . . . . . . 1

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Part I Thermodynamics

2 Within and Without: Systems and Surroundings . . . . . . . . . . . . . . . . . . . 7

2.1 Isolated Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.2 Closed Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.3 Open Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4 Life is an Open System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

3 Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

4 Matter and Energy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

4.1 Matter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
4.2 Energy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
4.3 Internal Energy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
4.4 Internal Energy (U) Exchanges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

5 Energy Accountability: Enthalpy (H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

5.1 The Chemical Reaction System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
5.2 Chemical (Standard) Enthalpy Exchanges . . . . . . . . . . . . . . . . . . . . . . 28
5.3 Chemical (Nonstandard) Enthalpy Exchange . . . . . . . . . . . . . . . . . . . . 31
Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

6 Energy Has Bias: Entropy (S) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

6.1 Second Laws of Thermodynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
6.2 Energy Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

xi
xii Contents

7 The Energy Exchange Gradient: Gibbs Energy (G) . . . . . . . . . . . . . . . . 43

7.1 ∆ G◦ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
7.2 Energy Unification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
7.3 ∆ G◦ : Closed Systems Under Standard Conditions . . . . . . . . . . . . . . . 47
7.4 ∆ G: Nonstandard Conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Part II Bioenergetics

8 Life’s Currency: ATP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

8.1 ATP: Structure and Content . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
8.2 ATP: Energy Exchange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
8.3 ATP: Turnover Efficiency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
8.4 ATP Utilization (Energy Demand) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
8.5 ATP Resynthesis (Energy Supply) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

9 Metabolism as an Energy-Exchange Device . . . . . . . . . . . . . . . . . . . . . . . 63

9.1 Metabolic Power: Force and Flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
9.2 Negative Entropy (?) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
9.3 Dynamics of a Metabolic Pathway . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
9.4 Intracellular Transport . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
9.5 Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
9.6 Exergy Synopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

10 Anaerobic Metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
10.1 A Brief History of Anaerobic Biochemistry . . . . . . . . . . . . . . . . . . . . . 75
10.2 The Glycolytic Gradient . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
10.3 Glycolytic Enthalpy and Entropy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
10.4 “High-Energy” Phosphate Buffering . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
10.5 Anaerobic “Speed” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

11 Aerobic Metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
11.1 Mitochondria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
11.2 Krebs Cycle: Gradient 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
11.3 Electron Transport Chain: Gradient 2 . . . . . . . . . . . . . . . . . . . . . . . . . . 91
11.4 Proton-Motive Force: Gradient 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
11.5 The Creatine Phosphate Shuttle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
11.6 ATP Tally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Contents xiii

Part III Metabolism

12 Aerobic Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99

12.1 Combustion, Respiration, and Heat . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
12.2 Thornton’s Law: Combustion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
12.3 Respiration and Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
12.4 Heat and Gas Exchange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
12.5 Aerobic Energy Expenditure as Heat and Entropy . . . . . . . . . . . . . . . 106
12.6 CO2 and O2 : RER = Aerobic and Anaerobic Energy Exchange . . . . 108
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110

13 Anaerobic Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111

13.1 The Oxygen Deficit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
13.2 Lactate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

14 Metabolic Energy Expenditure at Rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123

14.1 Measuring Energy Expenditure: Calorimetry . . . . . . . . . . . . . . . . . . . . 123
14.2 The Energy Expenditure of Rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
14.3 Eating and Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
14.4 Pregnancy and Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133

15 Metabolic Energy Expenditure of Activity (Work and Exercise) . . . . . 137

15.1 Rate vs. Capacity vs. METs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
15.2 Muscle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
15.3 Work and Energy Expenditure Relationships . . . . . . . . . . . . . . . . . . . . 143
15.4 Glycolytic vs. Respiratory Efficiency . . . . . . . . . . . . . . . . . . . . . . . . . . 145
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

16 Total Energy Expenditure of Exercise and Recovery . . . . . . . . . . . . . . . 149

16.1 Aerobic Exercise Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . 149
16.2 Anaerobic Exercise Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . 151
16.3 Aerobic Recovery Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . 153
16.4 Dismissing the Oxygen Debt Hypothesis . . . . . . . . . . . . . . . . . . . . . . . 155
16.5 Total Energy Expenditure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
16.6 Weight Loss: Low vs. High Intensity Activity . . . . . . . . . . . . . . . . . . . 158
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Chapter 1
Introduction: Thermodynamics, Bioenergetics,
Metabolism

Some things feel hot, some cold. This sense has meaning. Heat comes and goes,
appearing and seemingly disappearing; thermodynamics studies thermal activity.
The measurement of heat helps portray fundamental aspects of matter and energy.
Originally devoted to uncovering the hows and whys of a steam engine’s ability to
convert heat into mechanical work, thermodynamics quickly expanded into the field
of chemistry . . . and more. We have known for many thousands of years, perhaps far
longer, that life and heat are related (1). Because of your association with thermal
energy, a proper examination of nutrition and exercise cannot be initiated without
addressing thermodynamics to some extent.
Matter and energy are distributed about the Universe and the Earth, and in similar
fashion throughout you and all your cells. These distributions are dynamic; energy
undergoes transfer, so too does matter, from one place to another. The transfer of
matter and energy can be spontaneous, or not. A gradient – a difference over a
distance (2) – can both promote and prevent the “flow” of energy and materials.
Gradients empower dynamics. Perhaps the most obvious example of a gradient is
founded in our planet’s topography where gravity is the driving force and the spon-
taneous direction of change is downhill; uphill gradients are said to be nonsponta-
neous (Fig. 1.1).
Up and down gradients can also be expressed in left and right formats. Chemistry
for example uses a left-to-right and right-to-left format to respectively describe the
direction of spontaneous and nonspontaneous chemical reactions:

High → Low

A left-to-right arrow signifies a downhill or spontaneous gradient.

High ← Low

A right-to-left arrow signifies an uphill or nonspontaneous gradient.

The secret to life is a gradient. Life uses a variety of gradients to facilitate matter
and energy exchanges. The cardiovascular system for example operates off a pres-
sure gradient measured in millimeters of mercury (mm Hg). Upon contraction the

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2 1 Introduction: Thermodynamics, Bioenergetics, Metabolism

High

Low

Fig. 1.1 A gradient is shown in the format of a ramp. If placed on this gradient, matter, in the form
of a rubber ball, would be transferred in the downhill direction, from uphill to downhill. Energy
too flows naturally in the downhill direction. Uphill transfer is nonspontaneous (in fact, energy is
required to “push” matter uphill)

heart creates high pressure within the arteries; within veins the pressure is much
lower. The result is blood flow, from high pressure to low, arteries to veins. Gases
too flow from areas of high-to-low pressure, the result of a partial pressure gradient
(also measured as mm Hg).
In addition to transfer from one place to another, matter and energy also undergo
conversion from one form to another. Survival is dependent on devices that favorably
enable matter and energy conversions. We understand steam engines, automobiles,
refrigerators, light bulbs, and the like, as devices or machines that convert matter
and energy from one form to another. An active metabolism – the biochemistry con-
tained within all your bodies cells – has been identified as the place(s) where life’s
energy and matter conversions take place. Energy transfer and energy conversion
are both collectively described in the context of energy exchange. Bioenergetics is
a continuing scientific narrative, a human attempt to identify the source of all en-
ergy and material exchanges (i.e., transfer and conversion) within and without life.
Biology, nutrition, physiology, chemistry, physics, engineering – bioenergetics is all
these and more.
The costs of living are defined by studying the conversion and transfer of matter
and energy (exchanges) within and without the boundaries that separate life from
its immediate surroundings. Energy and matter exchanges within cells are the re-
sult of a working metabolism, often considered the biochemistry of life. All life
forms share something in common: continuous matter and energy intake from, and
References 3

matter and energy expenditure to, the environment. By measuring energy intake in
the form of food and drink, for example, a nutritionist may gain an understand-
ing of the energy and material needs of survival, or in the case of an athlete, the
needs of training and competing. A measurement or estimate of energy expenditure
can likewise be used in an attempt to identify and account for the costs of living
and working. Sometimes the two reciprocal movements – intake and expenditure –
through a boundary need to be considered for a more complete analysis of the ex-
changes made between a living organism and its surroundings. One example of this
is the quantification of oxygen uptake taken from and carbon dioxide given-off to
the environment. Indeed, the measurement of metabolic gas exchange continues to
be a powerful tool in providing an estimate of the costs of living, training, and com-
peting.

References

1. Mendelsohn, E. Heat and life. Cambridge, MA: Harvard University Press, 1964.
2. Schneider ED, Sagan D. Into the cool: energy flow, thermodynamics and life. Chicago:
University of Chicago Press, 2005.
Chapter 2
Within and Without: Systems and
Surroundings

A specified region or space that is surrounded by a boundary is called a system. The

surroundings reside outside of the system. Sometimes the boundary in mind between
system and surroundings is not explicit, but, real or imagined, it must be carefully
defined. Take, for example, our solar system. That is a big system to be sure, whose
boundaries appear more subjective than objective. Yet our solar system is soundly
understood in the context of what it contains: the sun, eight planets, along with Pluto,
various other moons, comets, asteroids, etc. On other scales, a single atom, a pro-
tein molecule, a chemical reaction, a linked-together series of biochemical reactions
(a metabolic pathway), a living cell, also each represent unique systems located
within a specified surrounding.
Life too shares a boundary with its surroundings. The human system resides
within the boundary of its skin; many types of exchanges take place with our en-
vironment: the consumption of food, the excretion of waste, oxygen uptake, car-
bon dioxide, and heat production are but a few examples. Nature often compli-
cates matter and energy exchanges. A hot muggy day for example may create a
life-threatening reversal of heat exchange for an exercising athlete, from rather than
to the environment.
How many cells are you made of? Each represents an independent system with
its external membrane allowing for a selective exchange of matter and energy with
the surroundings (Fig. 2.1). Cells are also highly compartmentalized within creating
further internal boundaries that contribute to the unequal distribution of interior mat-
ter and energy. Systems and surroundings interact or do not interact depending on
the type of border involved. External and internal cell borders create and exploit en-
ergy and matter gradients by separating a system from the immediate surroundings.
Boundaries enable gradients, gradients empower life.

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DOI 10.1007/978-1-60327-383-1


c Humana Press, a part of Springer Science+Business Media, LLC 2008